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the development of the mammalian retina, a diverse range of cell types is generated from a pool of multipotent retinal progenitor cells. Among these cell types, photoreceptors account for over 70% of all cells in the retina. In vertebrates, there are two classes of photoreceptors, rods and cones. Rods are sensors of dim light, while cones function in bright light and are responsible for color vision. Phototransduction, a series of signal amplifications detecting a single photon of light, is initiated by the capture of light with 11-cis-retinal, a chromophore bound by the opsin proteins: rhodopsin in rods and cone opsins in cones. The proteins that carry out phototransduction are located in an elaborate and highly specialized membranous structure, the outer segment. The outer segment appears to be relatively fragile, degenerating in response to many environmental and/or genetic perturbations. In the rodent retina, the production of specific cell types during development progresses in a general order [1,2]. Rod photoreceptor generation peaks around the time of birth. Cone photoreceptors, ganglion cells, horizontal cells and amacrine cells are generated earlier, while Müller glia and bipolar cells are generated later. This production of different cell types at different times appears to derive from differences in the intrinsic properties of progenitor cells involved in the transcription or chromatin modification. Recent studies identified several important transcription factors of photoreceptor development [3-7]. Two Otx family homeobox genes, Otx2 and Crx, play essential roles in early cell fate determination and terminal differentiation of photoreceptors [3,8,9]. In the absence of Otx2 function, differentiating photoreceptor cells are converted to amacrine-like neurons [9]. Crx, a downstream target of Otx2, controls the transcription of various photoreceptor cell-specific genes and is essential for the formation of outer segments, synaptic terminals, and phototransduction pathways [8,10]. Crx transcripts begin to be expressed in developing photoreceptors at embryonic day 12.5 (E12.5) in the mouse and a strong upregulation of Crx transcription is apparent across the differentiating photoreceptors at postnatal day 6 (P6). Photoreceptor cells in the Crx knockout (KO) mice exhibit a dramatic reduction of many photoreceptor molecules including visual pigments and develop neither photoreceptor outer segments nor a synaptic terminus [8,10]. In addition, mutations of human CRX have been demonstrated to be associated with three types of photoreceptor diseases: autosomal dominant cone-rod dystrophy 2, autosomal dominant-type retinitis pigmentosa, and Leber's congenital amaurosis (LCA) [11-14]. While Otx2 and Crx control general photoreceptor development, three other transcription factors, TRβ2, Nrl, and Nr2e3 regulate the specification of photoreceptor cell types [4,5,7]."}

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the development of the mammalian retina, a diverse range of cell types is generated from a pool of multipotent retinal progenitor cells. Among these cell types, photoreceptors account for over 70% of all cells in the retina. In vertebrates, there are two classes of photoreceptors, rods and cones. Rods are sensors of dim light, while cones function in bright light and are responsible for color vision. Phototransduction, a series of signal amplifications detecting a single photon of light, is initiated by the capture of light with 11-cis-retinal, a chromophore bound by the opsin proteins: rhodopsin in rods and cone opsins in cones. The proteins that carry out phototransduction are located in an elaborate and highly specialized membranous structure, the outer segment. The outer segment appears to be relatively fragile, degenerating in response to many environmental and/or genetic perturbations. In the rodent retina, the production of specific cell types during development progresses in a general order [1,2]. Rod photoreceptor generation peaks around the time of birth. Cone photoreceptors, ganglion cells, horizontal cells and amacrine cells are generated earlier, while Müller glia and bipolar cells are generated later. This production of different cell types at different times appears to derive from differences in the intrinsic properties of progenitor cells involved in the transcription or chromatin modification. Recent studies identified several important transcription factors of photoreceptor development [3-7]. Two Otx family homeobox genes, Otx2 and Crx, play essential roles in early cell fate determination and terminal differentiation of photoreceptors [3,8,9]. In the absence of Otx2 function, differentiating photoreceptor cells are converted to amacrine-like neurons [9]. Crx, a downstream target of Otx2, controls the transcription of various photoreceptor cell-specific genes and is essential for the formation of outer segments, synaptic terminals, and phototransduction pathways [8,10]. Crx transcripts begin to be expressed in developing photoreceptors at embryonic day 12.5 (E12.5) in the mouse and a strong upregulation of Crx transcription is apparent across the differentiating photoreceptors at postnatal day 6 (P6). Photoreceptor cells in the Crx knockout (KO) mice exhibit a dramatic reduction of many photoreceptor molecules including visual pigments and develop neither photoreceptor outer segments nor a synaptic terminus [8,10]. In addition, mutations of human CRX have been demonstrated to be associated with three types of photoreceptor diseases: autosomal dominant cone-rod dystrophy 2, autosomal dominant-type retinitis pigmentosa, and Leber's congenital amaurosis (LCA) [11-14]. While Otx2 and Crx control general photoreceptor development, three other transcription factors, TRβ2, Nrl, and Nr2e3 regulate the specification of photoreceptor cell types [4,5,7]."}

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the development of the mammalian retina, a diverse range of cell types is generated from a pool of multipotent retinal progenitor cells. Among these cell types, photoreceptors account for over 70% of all cells in the retina. In vertebrates, there are two classes of photoreceptors, rods and cones. Rods are sensors of dim light, while cones function in bright light and are responsible for color vision. Phototransduction, a series of signal amplifications detecting a single photon of light, is initiated by the capture of light with 11-cis-retinal, a chromophore bound by the opsin proteins: rhodopsin in rods and cone opsins in cones. The proteins that carry out phototransduction are located in an elaborate and highly specialized membranous structure, the outer segment. The outer segment appears to be relatively fragile, degenerating in response to many environmental and/or genetic perturbations. In the rodent retina, the production of specific cell types during development progresses in a general order [1,2]. Rod photoreceptor generation peaks around the time of birth. Cone photoreceptors, ganglion cells, horizontal cells and amacrine cells are generated earlier, while Müller glia and bipolar cells are generated later. This production of different cell types at different times appears to derive from differences in the intrinsic properties of progenitor cells involved in the transcription or chromatin modification. Recent studies identified several important transcription factors of photoreceptor development [3-7]. Two Otx family homeobox genes, Otx2 and Crx, play essential roles in early cell fate determination and terminal differentiation of photoreceptors [3,8,9]. In the absence of Otx2 function, differentiating photoreceptor cells are converted to amacrine-like neurons [9]. Crx, a downstream target of Otx2, controls the transcription of various photoreceptor cell-specific genes and is essential for the formation of outer segments, synaptic terminals, and phototransduction pathways [8,10]. Crx transcripts begin to be expressed in developing photoreceptors at embryonic day 12.5 (E12.5) in the mouse and a strong upregulation of Crx transcription is apparent across the differentiating photoreceptors at postnatal day 6 (P6). Photoreceptor cells in the Crx knockout (KO) mice exhibit a dramatic reduction of many photoreceptor molecules including visual pigments and develop neither photoreceptor outer segments nor a synaptic terminus [8,10]. In addition, mutations of human CRX have been demonstrated to be associated with three types of photoreceptor diseases: autosomal dominant cone-rod dystrophy 2, autosomal dominant-type retinitis pigmentosa, and Leber's congenital amaurosis (LCA) [11-14]. While Otx2 and Crx control general photoreceptor development, three other transcription factors, TRβ2, Nrl, and Nr2e3 regulate the specification of photoreceptor cell types [4,5,7]."}